JOURNAL OF ANIMAL BEHAVIOR

VOL. 4 SEPTEMBER-OCTOBER, 1914 No. 5 A STUDY OF SEXUAL TENDENCIES IN MONKEYS AND BABOONS
G. V. HAMILTON Montecito, California

In spite of the considerable advance that has been made in our knowledge of sexual life since the appearance of Freud's (1) Drei Abhandlungen zur Sexualtheorie, we still lack that knowledge of Infra-human sexual life without which we may scarcely hope to arrive at adequately comprehensive conceptions of abnormal human sexual behavior. For example, the possibility that the types of sexual behavior to which the term " perverted " is usually applied may be of normal manifestation and biologically appropriate somewhere in the phyletic scale has not been sufficiently explored. Homosexual tendencies come to frequent expression in adolescent boys and girls, thereby presenting to the mental hygienist a problem, >the solution of which awaits, first of all, biological knowledge of homosexuality which only the behaviorist can supply. It is unnecessary to multiply examples in illustration of the fact that both the theoretical interests of the science of behavior and the practical needs of what we may regard as a group of applied sciences of human behavior (viz., mental hygiene, criminology, psychopathology) place upon the animal behaviorist an obligation to lay the necessary foundations for a scientific and thoroughly comprehensive. investigation~~of sexual life. The above considerations, of which I have been almost daily reminded by clinical contacts with human sexual problems, have led me to formulate the following problems in animal behavior: (1) Are there any types of infra-human primate behavior

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which cannot be regarded as expressions of a tendency to seek sexual satisfaction, but which have the essential objective characteristics of sexual activity ? (2) Do such sexual reaction-types as homosexual intercourse, efforts to copulate with non-primate animals and masturbation normally occur among any of the primates, and if so, what is their biological significance ? It is always a difficult matter to collect scientific data which shall be specifically relevant to behavior problems when such problems do not lend themselves to strictly experimental methods of investigation. Even under carefully prearranged experimental conditions one cannot always be sure that adventitious stimuli may not have played a part in bringing about a given response. This uncertainty is much greater when the animal subject is either at large with his fellows or confined with them in a cage of sufficient size to allow a reasonable approximation to natural conditions. Under such circumstances the best that one can do is to supply a set of conditions which are apt to lead to a fairly definite and uncomplicated development of the desired situation. The monkey's marked variability of response presents a further difficulty, for in seeking to identify a definite situation-response sequence the observer is called upon to distinguish activities that are essential components of a given reaction-type from purely fortuitous activities. For example, in a given^case a monkey's manipulation of his genitalia may be nothing more than reflex scratching of a momentarily irritated area, and not at all a part of his response to the situation of which the observer wishes to determine the reactive value. In many cases I have been unable to ascertain the essential components of a given reaction-type until prolonged contact with my subjects has enabled me to predict with reasonable certainty that whenever the appropriate situation developed a sequence of activities composed of such and such members would be manifested. A difficulty of another kind is encountered when one seeks to present results that have been obtained by non-experimental methods. Such results can seldom be indicated by tables of figures, habit-formation curves, etc., because reaction to " natural " situations are usually complexes of activities which call for detailed description. When, as in the present instance,

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the observer arrives at conclusions which have no value unless they are found to be justified by the facts upon which they are based, and when the presentation of all the facts involved would require the. space of several printed volumes, some method of abridgment must be adopted. An abridged journal is not apt to be satisfactory where an extensive program of observation has been followed unless, as rarely happens, the most convenient and logical order in which the facts can be presented coincides with the order of their occurrence. I have followed the order of presentation outlined below for the sake of effecting a satisfactory abridgment without resorting to the awkward expedient of publishing extracts from my note books in journal form: I. II. III. List of subjects. Description of environmental conditions. A list of the types of situations that were arranged by the observer or encountered by the subjects in consequence of their spontaneous activities; and under each description of a typical situation one or more detailed descriptions of typical responses thereto. IV. Classification of reactions as expressions of reactive tendencies.

I. DESCRIPTION OF SUBJECTS The estimated or known age of each subject is given for January, 1914. Inability to identify an animal as to species is indicated by a dash after the generic name. The " pet " name of each animal is given to facilitate the reader's identification of subjects in subsequent descriptions of behavior. An animal's sex is indicated by its laboratory numbereven numbers for males, odd for females. In reporting the behavior of an animal the first reference to it will include, in the order given, its laboratory number, pet name and initial letters of the genus and species to which it belongs. E.g., " 7-Becky-M-r " refers to Monkey 7 of the list, and indicates that she is a female M. rhesus. Monkey 1. Bridget. M. rhesus. Adult. Monkey 2. Mike. M. adult. About 1/4 larger than adult male M. rhesus. Fur grey and luxuriant. Tail about 8 centimeters long and furred to the tip. Body thick, face broad. Readily identifiable as a macaque.

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Monkey 3. Kate. M. rhesus. Adult. Mother of Monkey 9. Monkey 4. Pat. M. rhesus. Adult. Vision defective. Monkey 5. Maud. M. rhesus. Young adult. Monkey 6. Jocko. M. cynomolgus. Adult. Monkey 7. Becky. M. rhesus. Adult. Mother of Monkeys 13 and 24. Monkey 8. Jimmy I. M. cynomolgus. Adult. Monkey 9. Gertie. M. cynomolgus-rhesus. Age, 3 years, 2 months. Daughter of monkeys 3 and 10. First pregnancy began September, 1913. Monkey 10. Timrny. M. cynomolgus. Adult. Father of Monkeys 9, 13 and 24. Baboon 11. Grace. Papio. Adult. A small black baboon, about 1-3 larger than an adult female M. rhesus. Tail absent, other anatomical features similar to those of typical members of genus Papio. Became pregnant when bred to male baboon (Monkey 12) but frequent previous copulation with male macaques was without result. Baboon 12. Sandy. Papio. Adult. Fur black over back and forehead, grey elsewhere. " Pig " tail, long muzzle. Legs and body much shorter and thicker and those of the chacma. A very powerful animal, almost equal to the chacma in weight. Monkey 13. Tiny. M. cynomolgus-rhesus. Age, 5 months. Daughter of Monkeys 7 and 10. Monkey 14. Jimmy II. M. cynomolgus. Young adult. Monkey 16. Sobke. M. rhesus. Young adult. Monkey 18. Baby. M. cynomolgus. About 1-2 adult size. Castrated before sexual maturity. Monkey 20. Chatters. M. cynomolgus. About 3-4 adult size. Castrated before sexual maturity. Monkey 22. Daddy. M. cynomolgus. Adult. Castrated date unknown. Monkey 26. Skirrel. M. cynomolgus. Adult. Monkey 28. Scotty. M. Young adult. Probably belongs to the cynomolgus group.
II. ENVIRONMENTAL CONDITIONS

My laboratory is in the midst of a live oak woods in Montecito, California, about five miles from Santa Barbara. Like all of

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Southern California, Montecito is not entirely free from frost, but the winters are so mild that when the orange and lemon growers far to the south of us are compelled to use artificial heat in their orchards to prevent damage by frost, our local growers find it unnecessary to take any precautions whatsoever against the cold. The climate here is therefore exceptionally mild, even for Southern California, and at no times seems to reduce the activities of macaquesexcept, of course, when rain drives them to shelter. My subjects have always been in excellent physical condition, the only deaths having been due to accident or to pathological causes that were operative at the time of an animal's purchase. In front of the laboratory is a quadrangular yard, 16.7 meters long by 7.4 meters wide. The laboratory encloses one end of this yard, and the cage one side. The other end and side of the yard is enclosed by a solid board fence which is 1.9 meters high. The entire enclosure is surrounded by live oak trees, and one tree is contained within the yard. The animals, when at large, could wander to an indefinite distance from the laboratory by passing from tree to tree, but they rarely wander out of sight of the yard. The cage is 6 meters high, 16.7 meters long and 1.8 meters wide. The front, top and upper half of the rear and ends are covered with wire netting, the meshes of which are 1.4 centimeters square. The lower half of the rear and ends is solidly boarded, to give stability to such a tall, narrow structure. The cage is subdivided into eleven compartments by partitions of which the lower one third is wood and the upper two thirds wire netting (1.4 cm. mesh). Within 72 centimeters of the top of each compartment is a horizontal shelf, 30 centimeters wide. Each compartment is also equipped with a sleeping box, a food drawer, and proper drainage for the concrete floor. A door at the rear gives access to the man who cleans the cage. An important accessory to the cage is a wooden alley, which extends along the entire rear of the structure, midway between the top and bottom. This alley is 75 centimeters high by 60 centimeters wide. Each compartment opens into the alley by means of a sliding door arrangement, which enables the observer to make it accessible to the occupants of one or more compartments, according to the demands of a given experiment.

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G. V. HAMILTON III. TYPES OF SITUATIONS AND OF RESPONSES THERETO

The outdoor conditions described above have enabled me to liberate most of the subjects in selected pairs or groups. 13Tiny-M-cr was too young, and 10-Timmy-M-c, 12-Sandy-Papio, 26-Skirrel-M-c and 28-Scotty M were too large to be safely liberated, but an excellent substitute for outdoor freedom was obtained for them by giving them access to the long alley and three or four of the cages. I have been able to form a fairly accurate estimate of the effects of cage-life on sexual behavior by comparing the activities of animals that have been at large for several months with the activities of those that have been continuously confined. At this point it may be said that, provided the macaque or baboon have a sufficiently wide range for pursuing and fleeing his enemies, playing with his fellows, etc., confinement will be found to have no perverting effects in this climate. In the following list of observations I have endeavored to spare the reader the inconvenience of being compelled to make frequent reference to the list of subjects by giving in each instance the age of the animals under discussion, rather than the date of the observation. Wherever reference is made to an animal as " sexually immature " it is to be understood, in the case of females, that menstruation has not yet appeared; males are regarded as sexually immature until they have assumed the characteristic strut that I have found to be coincident _in appearance with seminal discharge on copulating with females. Sexual maturity, in this sense, antedates the attainment of adult size by at least a year. Situation 1. Male and female separated from one another (but not from fellows of the opposite sex) for at least one week, then given access to one another. Observation 1. 10-Timmy-M-c and 5-Maud-M-r, both sexually mature, occupied separate cages, each with a mate, for several weeks. Each animal had been copulating freely. They were then given access to one another, their respective mates having been tolled into an empty cage and confined therein. Timmy rushed from the alley into Maud's cage as soon as her door was opened. She observed his approach from her shelf, and as he ascended toward her, smacking his lips, she, too, smacked her lips. As soon as the male clambered upon the shelf the female

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assumed the sexual position, viz.: Hind legs fully extended to an almost vertical position; forelegs sharply flexed; tail erect; body inclined forward and downward from the hips; head sufficiently extended and rotated to enable the female to direct her gaze upward and backward. The male grasped the female at the angles formed by the juncture of hips and body with a hand on either side, and in mounting her, he clasped her legs just above the knees with his feet. He leaned forward and downward during copulation, smacking his lips violently. The female seemed to invite contact with his mouth, for she persistently thrust her smacking lips towards the male, until he leaned still further downward and touched her lips with his own. Shortly before copulation ceased, the male uttered a succession of shrill little cries, and greatly increased the vigor of his copulatory movements. As soon as he dismounted the female, he took her tail in one hand and elevated it, then with his free hand examined her vaginal labia, at the same time closely inspecting them with nose and eyes. Then he lay down and the female examined his fur " flea-hunting." "Observation 2. 11-Grace-Papio (sexually mature), 14-JimmyM-c (sexually mature), the three eunuchs and a number of immature monkeys had been at large for several weeks. Grace had freely copulated with all of the males, including the eunuchs. 16-Sobke-M-r (sexually mature) had occupied a cage with a mature female for several weeks. He was now liberated. As soon as he and Grace caught sight of one another they began smacking their lips. Their subsequent behavior was essentially similar to that of the animals described in observation 1. Situation 2. Male and female after several days' access to one another, during which copulation had been of frequent occurrence. Observation 3. After Timmy and Maud had been together one week (date of this observation is one week later than that of observation 1, q.v.) Timmy was offered a loquat. He descended from his shelf, leaving Maud still perched upon the shelf. As soon as he had reached the floor, and had received and eaten the loquat, he began to stalk his mate, slowly ascending with tail erect, gaze fixed upon the female, and head jerked forward now and then as if to threaten violence to Maud. Such behavior often precedes an attack upon another male. When he finally

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attained the shelf he copulated with Maud in the manner described in observation 1. After a few minutes he again mounted her, but this time the female merely arose from a crouching to a quadrupedal standing position, without displaying signs of sexual excitement. There was no smacking of lips by either animal, and the desultory efforts of the male to copulate terminated as soon as he entered the female and made two or three strokes. Both animals sat inactive for five minutes, then the male drew the female's hind quarters towards him. He started to mount, then desisted and began to inspect her genitalia with eyes, nose, mouth and fingers. After two or three minutes of this behavior he copulated, both animals displaying sexual excitement. During the next half hour there was an alternation of perfunctory attempts at copulation with more vigorous attempts, the latter being immediately preceded by examinations of the female's genitalia on the part of the male. Observation 4. 14-Jimmy II-M-c and 9-Gertie-M-cr (both sexually mature young adults) had been at large together in company with a number of other males and females for several weeks. They had copulated a number of times each day. Their behavior during a half hour when they were perched together upon the roof of the laboratory consisted in the same alternation of perfunctory attempts at copulation with more vigorous attempts, the latter being immediately preceded by examination of the female's genitalia on the part of the male. Jimmy II is more apt to follow me about when- "he Is at large than to stay with the tribe, and this may account for the fact that he manifested the behavior just described less frequently than did Timmy. 26-Skirrel-M-c manifests such behavior more frequently than does Jimmy II and less frequently than does Timmy. The mature rhesus male (16-Sobke-M-r) also inspects the female's genitalia after a perfunctory attempt at copulation and just before a vigorous attempt, but he is more apt to increase his sexual excitement by chasing and biting the female. All of my mature males, with the exception of the eunuchs and the baboon, at times chase and bite their mates in apparent preparation for copulation, but only after prolonged access to them. Situation 3. Two males, confined in separate cages with mates, are caused to exchange mates.

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Observation 5. 16-Sobke-M-r and 3-Kate-M-r (both sexuallymature) had occupied a cage adjacent to that occupied by 10-Timmy-M-c and 11-Grace-Papio (both sexually mature). The males had ceased to copulate vigorously with their mates without first examining their genitalia or chasing and biting them. When Timmy was now separated from Grace and put into Kate's cageSobke having previously been removed-he dashed up the side of the cage toward Kate and copulated with her vigorously without any preliminary attentions to her. The female displayed sexual excitement, although she had ceased to do so when copulating with Sobke unless he first examined her genitalia or chased and bit her. As soon as this observation was completed, Sobke was allowed to enter Grace's cage. He, like Timmy, had ceased to copulate vigorously unless he first examined or chased the female. But as soon as he was given access to Grace he displayed great sexual excitement, and copulated vigorously, without having engaged in any preliminary activities conducive to sexual excitement. After a few days the behavior of the newly mated couples toward one another was essentially similar to their behavior toward their former mates. When they were restored to their former mates, vigorous copulation again ensued without preliminary examining and chasing on the part of the males. Situation 4. Female taken from her mate and given to a male in an adjacent cage, her mate being left alone in his cage. Observation 6. 10-Timmy-M-c and 11-Grace-Papio had occupied a cage together for several weeks, and 26-Skirrel-M-c had been alone in an adjacent cage. All subjects were sexually mature. Grace was taken from Timmy and given to Skirrel. As soon as Skirrel began to copulate with Grace on the shelf of their cage Timmy, who was on his shelf and therefore in very close proximity to his former mate and to Skirrel, began to rage at Skirrel through the netting. Skirrel paid no attention to his baffled enemy, although under ordinary circumstances he would have come to the netting and made a hostile demonstration, these two animals being enemies. Timmy continued to rage at Skirrel until the latter had ceased copulation and had assumed a semi-recumbent position on the shelf. Then Timmy went about a foot from the netting and assumed the female position

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for copulation, smacking his lips as if to invite copulation with SkirrelGrace had descended to the floor in quest of food. As soon as Skirrel approached the netting Timmy whirled and thrust his forefinger through the netting into Skirrel's eye. Situation 4. Male taken from his mate and given to a female in an adjacent cage, the first female being left alone. Observation 7. 10-Timmy-M-c and 11-Grace-Papio had occupied a cage together for a week, and 7-Becky-M-r occupied an adjacent cage. (All subjects sexually mature.) Timmy was taken from Grace and given to Becky. Timmy copulated with his new mate on the shelf. Grace, now alone and on her shelf, barked angrily and tried to get at Becky through the netting. Timmy was tolled to the floor, leaving Becky and Grace on their shelves, and separated only by the netting. Grace tried to engage Becky in a combat through the netting, but Becky withdrew to the opposite end of the shelf. Then Grace began to smack her lips, and assumed the sexual position, although the male was not in sight. Becky approached the netting, whereupon Grace whirled (she had thrust her hind quarters against the netting) and poked at Becky's face through the netting. Situation 5. Female at large with band containing dominant male and a sexually mature smaller male. Female detected by dominant male while she is copulating with the smaller male. Observation 8. 11-Grace-Papio, 16-Sobke-M-r and 14Jimmy II-M-c (all three animals sexually mature) were at large with a band containing eight other monkeys. Sobk dominated the band, and Jimmy II had not yet learned to dominate Grace. Sobke and Grace frequently copulated, but at times Grace tolled Jimmy II to a place secluded from Sobke and invited him to copulation. The male baboon's sexual organ being much larger than the male macaque's and of different formation, Grace's sexual hunger seems to be insatiable when she is denied access to the male baboon. Sobke was perched in a tree outside the yard, and Grace and Jimmy were in the yard, where I was seated making observations. The latter two animals were chattering at me, soliciting food, when Grace began to smack her lips and assumed the sexual position in invitation to Jimmy II. He mounted her, and copulation was well in progress when Grace and I heard a noise in the tree above us. We saw Sobke approaching, croaking angrily

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and evidently bent on an attack. Grace gave her hind quarters a quick jerk, dismounting Jimmy II. She then began to bark angrily and dashed at Jimmy II, who fled up the side of the cage to escape her attack. Sobke joined in the chase. When Jimmy II made good his escape Grace invited Sobke to copulation, thus terminating the episode. Observation 9. 1-Bridget-M-r, 4-Pat-M-r, and 2-Mike-M (sexually mature but not full grown) had been at. large several months. Pat was the dominant male and Bridget dominated Mike. During Pat's absence in a neighboring tree Bridget invited Mike to copulation. Pat descended upon them before the act was completed, and Bridget dismounted Mike with a great show of anger and led Pat in pursuit of Mike. This and the above episode may be regarded as characteristic of any monkey when in danger of an attack by a larger one. I have often made such observations as the following one: Two small monkeys were eating in close and peaceful proximity to one another. A larger monkey approached with threatening croaks, obviously intent on an attack. The monkey nearest the approaching enemy quickly turned upon his small companion, and by screaming at him angrily enticed the large monkey to join him in pursuit of his innocent victim. Such behavior is of daily occurrence among monkeys when they are at large. Situation 6. Female and five males having access to the alley and four cages. One of the males is sexually mature, one is sexually immature and three are eunuchs. A male baboon, many times larger than any of these animals, is given access to them: Observation 10. 11-Grace-Papio (sexually mature), 14Jimmy-II-M-c (sexually mature), 24-Mutt-M-cr (sexually immature) and the three eunuchs (one adult size and two smaller than adult size) had had access to the alley and four of the cages for a week. 12-Sandy-Papio (sexually mature and of great size as compared with the other five monkeys) was then allowed to enter the alley. Before Sandy's admission to the band Jimmy II, who had previously allowed Grace to dominate him, dominated the female and the four males. Grace copulated freely with him, but whenever she encountered any of the smaller monkeys in the alley during one of Jimmy II's frequent absences in one of

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the cages, she would copulate with them. I had frequently observed her surrounded by Mutt and the three eunuchs, copulating with one after another. The eunuchs were more apt to indulge in homosexual copulation than in copulation with the female, but they were not infrequently observed cohabiting, rather perfunctorily, with the female. For several days after Sandy's admission to the band the three eunuchs screamed with rage whenever he copulated with Grace. Jimmy and Mutt made no demonstration whatsoever on these occasions. The eunuchs' behavior is all the more remarkable in that, although they are ordinarily very timid, and especially afraid of Sandy, they would pull his fur when they found him copulating with Grace. Situation 7. Several males and females at large. Number of each sex sufficient to enable each animal to have frequent natural sexual relation. Observation 11. 16-Sobke-M-r (sexually mature), Jimmy IIM-c (sexually mature), 24-Mutt-M-cr (sexually immature), 22Daddy-M-c (adult eunuch), 20-Chatters-M-c (immature eunuch), 18-Baby-M-c (immature eunuch) and the following sexually mature females had been at large for several weeks: 11-GracePapio, 3-Kate-M-r, 7-Becky-M-r, 5-Maud-M-r and 9-GertieM-cr. They wandered freely about the laboratory yard, through the trees and over the tops of the laboratory and the cage. During this time I was able to make a large number*, of observations, which will be summarized here, rather than given separately and in detail. I found that, although Sobke dominated the band, and could therefore have as frequent sexual relations with any of the females as he desired, he, like all the other males, indulged in homosexual relations. His case is especially interesting, since he was acquired when quite small, and has spent much of his time at large, in company with both males and females. I cannot recall any time when he was confined without at least one female companion. Nevertheless, during the time now under discussion he would approach a smaller male, smack his lips and mount the object of his homosexual desires with evidence of considerable sexual excitement. He copulated with the females more frequently than with the males, and his homosexual behavior was of decidedly less frequent occurrence several months after he arrived at sexual maturity than it was during his sexual

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immaturity. The same kind of observations were made with reference to Jimmy II: even after he became sexually mature, and 'had free, outdoor access to females, he would use male monkeys as he used females, but this behavior became much less frequent after sexual maturity than it was during sexual immaturity. The eunuchs and the immature male (Mutt) were apt to detach themselves from the other members of the band, and to engage in such activities as mock fights, flea-hunts, races through the trees, etc. After a period of play they would huddle together, with arms around one another. Much smacking of lips would follow, and then they would copulate with one another, first one animal, then another, assuming the female position. During the time that these eleven monkeys were at large I was never able to observe a sexually mature (uncastrated) Monkey assume the female position for copulation with a weaker fellow. Sobke, for example, would use Jimmy II as he would use a female, but would not assume the female position for Jimmy II. On the other hand, even the adult eunuch would assume the female position for Mutt and Baby, both of whom were much smaller and weaker than he. Sobke was always larger than Jimmy II, but before these two animals arrived at sexual maturity one was as apt to assume the female position during homosexual relations as was the other. The only instance of homosexual relations between females that I could attribute to sexual desire on the part of both animals occurred when I allowed Kate to join the band. She had not had access to her daughter, Gertie, for more than a year, but as soon as she and Gertie met outdoors they rushed into an embrace, following which Gertie assumed the female position and Kate mounted her daughter. The mother made male copulative movements, and both animals smacked their lips and displayed marked excitement. This behavior was never repeated, although the two animals were at large together for several weeks. Situation 8. Mature male and immature male, having formed a friendship for one another, are separated. Observation 12. 14-Jimmy II-M-c (sexually mature) and 24-Mutt-M-cr (sexually immature) had been with a band consisting of females and other males. They all had access to the alley and four of the cages. Although Jimmy II cohabited

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with the females and the eunuchs, he and Mutt were usually seen perched upon a shelf together. Flea-hunting and affectionate embraces usually terminated in homosexual relations, Mutt always assuming the female position. After they had been together two weeks Jimmy II was removed from the band and placed in a cage with 7-Becky-M-r for one week. During this week of confinement with Becky, Jimmy II copulated with her freely. . At the end of the week he was allowed to rejoin the band. As soon as he and Mutt caught sight of one another they rushed into an embrace, smacking their lips and making crooning sounds. Both animals quickly manifested marked sexual excitment, and when Jimmy mounted Mutt the latter even rotated his head to bring his lips in contact with those of his friend. I have found that friendships between immature males and mature males are of frequent occurrence, and that they seem to have a sexual basis. Situation 9. Male, occupying cage with female, is supplied with a nonprimate companion, the female not being removed. Observation 13. 6-Jocko-M-c and 3-Kate-M-r, both sexually mature, occupied the same cage. A 4-foot gopher snake was thrown into their cage. Kate took refuge on her shelf, but Jocko cautiously approached the snake, lying stunned and almost motionless upon the floor. Jocko finally mustered sufficient courage to lift the snake's tail from the floor and inspect its anal orifice. He then bestrode the snake, seized its jaody in his two hands, and endeavored to copulate with it. When the snake began to squirm the monkey shrieked and fled to the top of the cage. But he made repeated attempts to approach the snake, which drove him away each time by blowing and thrusting out its tongue. Observation 14. Jocko, still occupying a cage with Kate, was supplied with a kitten, a puppy and a fox in succession. He attempted sexual relations with each of them after he had inspected their genitalia. Observation 15. A fox was given to a band of monkeys, including females and sexually mature 16-Sobke-M-r and 14-Jimmy IIM-c. Both males made frequent inspection of the fox's genitalia (the fox was a male) and attempted to copulate with it. 10Timmy-M-c manifested similar behavior when the fox was given to him, but he finally killed it.

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Situation 10. Females at large in a band containing males are given access to a male dog. Observation 16. 5-Maud-M-r and 3-Kate-M-r, both sexually mature, were at large with a band containing males. A small mongrel dog visited the yard now and then. On his approach all of the monkeys would take to the trees and chatter threateningly. But Maud finally descended to the ground and assumed the female position for copulation. The dog mounted her, dogfashion, and partially entered her. She displayed marked sexual excitement, and ever thereafter would descend to the ground and copulate with the dog whenever he entered the yard. Kate would descend to the ground and assume the sexual position, but would flee before the dog could embrace her. Maud finally offered herself to a strange dog, and the animal bit off her arm. Since then all of the monkeys have shown hostility to dogs. Situation 11. Male monkey in contact with human. Observation 17. 8-Jimmy I-M-c fsexually mature), while at large with a female, discovered a human infant lying in a hammock outdoors. The infant was too young to caress the monkey, but Jimmy I promptly endeavored to copulate with the infant. Observation 18 14-Jimmy II-M-r invariably attempts to copulate with my hand whenever I stroke him, in spite of the fact that he has been repeatedly cuffed for it, and has never been encouraged in such behavior. Situation 12. Mature male, after a week of confinement without companions. Observation 19. Of all my male monkeys only 6-Jocko-M-r (mature) has been observed to masturbate. After a few days confinement he would masturbate and eat part of his semen. I have reason to believe that he lived under unnatural conditions for many years before I acquired him In view of this fact that not one of seven sexually mature monkeys masturbated after several weeks of isolation under conditions that favored a fairly healthy mental and physical life (close proximity to other monkeys, large cage, warm climate), I am inclined to believe that masturbation is not of normal occurrence among monkeys. Situation 13. Mature female, after a week of confinement without a companion. Observation 20. 3-Kate-M-r, after a week of isolation, smacked her lips and assumed the sexual position whenever the male in

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t h e adjacent cage approached. A reported case of masturbation b y this animal proved t o be mere reflex scratching of t h e genitalia, which were irritated b y partially dried menstrual discharge. K a t e would assume t h e sexual position whenever I approached her cage, and when a ranch h a n d voided urine in her presence she displayed marked sexual excitement, smacking her lips a n d croaking invitingly. Observation 21. 5-Maud-M-r, after a week of isolation behaved in essentially t h e same manner as did K a t e in observation 20. Situation 14. Male, threatened b y larger male and unat* *",o U/V;> escape. Observation 22. When 12-Sandy-Papio' (mature) was '-^-H admitted to the alley band, of which 24-Mutt-M-cr (immature, was a member, all of the b a n d excepting Grace fled. S a n d y 0 will be remembered, is a very large baboon, a n d M u t t is1 immature rhesus, hence very tiny in comparison with Saf r ''j\ W h e n t h e b a n d scattered on t h e approach of the baboon, JN&Vt darted into a cage and crouched in a corner, on the floor. S a J "y followed him, leering a t him threateningly. M u t t squealed "tr> terror, a n d looked about him as if seeking an avenue of esr pe. Finding none, he assumed the female sexual position, b u t ? ly approached, now displaying only friendliness a n d sexual excitement, t h e little monkey darted between his legs a n d escaped. Observation 23. 28-Scotty-M occupied a cage-, adjacent t o t h a t occupied b y 10-Timmy-M-c. Scotty is sexually mi t e, b u t n o t yet of adult size. H e is very timid and Timrr . a savage adult. T i m m y forced the netting partition loo and entered Scotty's cage. Scotty fled in terror, shriekii and gnashing his teeth. W h e n T i m m y finally caught him an-' gan t o bite him Scotty assumed t h e female sexual position, ;eby effecting a temporary escape, since this diverted Timrr sufficiently to enable his victim t o d a r t past him. Situation 15. Female threatened b y larger female and i able to escape. Observation 24. 11-Grace-Papio and 9-Gertie-M-vDOth sexually mature) were brought together in the alley. oe a t once attacked Gertie and began t o bite her. Gertie ass^ the sexual position, and was mounted b y t h e larger f e m a k .race manifested no sexual excitement, and m a d e no copulat nove-

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ments. In a moment she returned to the attack, and Gertie again assumed the sexual position, but this time eluded the baboon and was allowed to escape to another cage. Qbservation 25. When 11-Grace-Papio (mature) was first admitted to an outdoor band of which 3-Kate-M-r (mature) was a member, she was attacked by Kate. For several days Kate was able to pounce upon the baboon's back, bite her and escape. But the baboon finally cornered her, whereupon Kate promptly assurned the sexual position, which did not avert the attack. I '? unnecessary to multiply instances of this kind, but it may bt osjaid that I have witnessed many episodes where a smaller i<~ "le endeavored to avert a larger female's attack by assuming t.4f^ sexual position, but that I have never known either animal ..,, ,display sexual excitement unless the aggressive female's Jientary mounting of her victim could be regarded as due to erotic impulse. The absence of such signs of sexual excite'it as smacking the lips and copulatory movements, and the cpndnued display of hostility on the part of the female while r is mounting the object of her attack render it unlikely that L " t a. mere act of mounting under these conditions occurs in respo^ 5e to sex hunger. On the other hand, the tendency of the wf t i female to offer herself to the aggressive female for copulatiou 3 clearly related to a desire to escape rather than to sexual 6 sire.. The next observation will tend to confirm this opinion. Situation 16. Female in a condition to render copulation int jceptable made accessible to a male. .%r.yation 26. A week after the birth of a full-term infant, wh\ ^was dead when first discovered, 9-Gertie-M-cr was made acce \1ible to 16-Sobke-M-r (both animals sexually mature). Bef ^'Cf,the birth of her infant Gertie had always been on good term. - " Ji Sobke, and had never displayed fear in his presence. But ijiien he approached her on this occasion, smacking his lips and "ager for copulation, she gnashed her teeth and squealed in fear.,L The male pursued her, and when she was cornered, Gertie asc< in. .1 the sexual position. When Sobke reared upon his hind \,gs Gertie darted past him. This episode was repeated sever imes, the female always effecting her escape before the mak aid copulate, and at no time displaying sexual excitement. Ay "ater she submitted to copulation with Sobke, and displaysv -xual excitement.

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Observation 27. 3-Kate-M-r (mature) was severely bitten by a dog while she was at large. Her tail was almost completely denuded of skin and fur, and her body was lacerated in places. She was put in a cage by herself for a week, then made accessible to 10-Timmy-M-c (mature). She was still weak and inclined to mope. Like Gertie in observation 26, she fled from the male, and assumed the sexual position only when she was cornered. The animals were separated as soon as it was apparent that her assumption of the sexual position was a purely defensive measure. Situation 17. Young sexually mature male attacked by sexual"y-mature female. Observation 28. 11-Grace-Papio (mature) was admitted to the cage occupied by 28-Scotty M. Scotty was sexually mature in the sense of being able to discharge the full sexual function, but had not yet attained adult size, and was very timid. Grace advanced toward him, barking and threatening an attack. When she began to bite him he assumed the female sexual position, whereupon Grace mounted him, "smacking her lips. She made no copulatory movements, and at once dismounted and assumed the female position. The male then copulated with her. Situation 18. Infant monkey in danger of attack. Observation 29. 13-Tiny-M-cr (age 5 | months) was momentarily abandoned by her mother (7-Becky-M-r) while the latter sat upon the sleeping box roof and fought the members of the alley band through the wire netting that separated tihem. Tiny was whimpering and running back and forth when 11-GracePapio dashed at her, barked, and clawed at the netting. Tiny's assumption of the sexual position had the appearance of a pure reflex, so quick was it, and so quickly followed by a retreat to her mother's arms. Observation 30. 24-Mutt-M-cr, when 6 months old, was observed to assume the female sexual position while his mother was absent from the shelf. He was whimpering and running back and forth on the shelf, when 10-Timmy-M-c (mature) threatened him from the other side of the netting. The infant assumed the female sexual position momentarily, then fled down the side of the cage to his mother. This kind of behavior may have been manifested earlier.

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IV. CLASSIFICATION OF SEXUAL REACTIONS AS EXPRESSIONS OF REACTIVE TENDENCIES

1. Tendencies to Seek Sexual Satisfaction. A. Male Tendencies (a) Tendency to engage in typical sexual intercourse with females. (b) Tendency to increase sexual excitement by preliminary examination of the female's genitalia, or by chasing and biting the female. (c) Tendency to use a younger or weaker male as a female. (d) Tendency to play the role of female to a copulating male. (e) Tendency to attempt copulation with non-primates and humans. (f) Tendency to masturbate (probably developed only under abnormal conditions). B. Female Tendencies (a) Tendency to engage in typical sexual intercourse with males. (b) Tendency to play the role of male to younger or weaker female. (c) Tendency to play the role of female to friendly female. (d) Tendency to solicit copulation with non-primates. Tendencies to Assume the Female Sexual Position as a Defensive Measure. A. Male Tendencies (a) Tendency to assume the female sexual position when attacked by a more powerful fellow of either sex. B. Female Tendencies (a) Tendency to assume the female sexual position when attacked by a more powerful fellow of either sex. 3. Tendencies to Seek to Lure an Enemy to Attack by Assuming the Female Sexual Position. A. Male Tendencies (a) Tendency to lure a male enemy to attack by assuming the female sexual position. 2.

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B.

Female Tendencies

(a) Tendency to lure a female enemy to attack by assuming the female sexual position. My analysis of the material from which the thirty observations recorded in the preceding pages were taken, at first inclined me toward a classification in which there would appear only the three general tendencies that appear in the above table, viz.: (1) A tendency to seek sexual satisfaction, (2) A tendency to assume the female sexual position as a defensive measure, (3) A tendency to lure enemies te attack by assuming the female sexual position. Had this scheme of classification been adopted, lists of the typical expressions of each of these three tendencies would have been given instead of the lists of specific tendencies that appear in the table. But the viewpoint from which I have come to regard animal and human behavior, taken inclusively, as material for a separate branch of natural science which may be made to serve as an important foundation for various applied sciences finally induced me to adopt the method of classification that appears above. According to my view, the behavior of an organism is the expression of reactive tendencies which have specific representation in its structure. Some of these tendencies have an inherent structural representation, such as, e.g., the tendency that finds expression in the kitten's spit and slap when it first encounters the dog-odorbehavior which may be observed in a kitten before its eyes are opened, and which earmot be attributed to the modifying effects of any previous experiences. Other tendencies owe their existence to two factors, viz.: (1) An inherent capacity for post-natal structural modifications by experience; these modifications by their appearance, add the tendencies of which they are the appropriate bases to the list of the organism's properties. (2) The operation of environmental influences that help to produce the necessary structural changes. Any habit-reaction may be regarded as the expression of an acquired tendency of this kind. My conception of behavior as reducible to expressions of specific reactive tendencies might easily lead to an endless multiplication of such tendencies to account for the apparently innumerable separate modes of organic activityespecially in view of the possibilities afforded by the extreme plasticity of the nervous systemif it did not include something more than the

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above generalizations. Lasurski (2), whose Hauptneigungen bear many points of resemblance to the reactive tendencies of my conception, avoids this danger by stopping short at the point at which he recognizes various relatively separate directions in which a given individual's activities may go in response to a limited number of principal inclinations. He regards these inclinations (tendencies) as artificial abstractions to which, nevertheless, human personality is most profitably and conveniently reducible. McDougall (3) calls attention to the fact that " The activities of each species are directed almost exclusively towards a small number of special endsreproduction, the securing of food, the escape from danger, the protection of the young, the violent destruction of whatever opposes these great tendencies, and a few others that differ from species to species." To this he adds: " The concentration of the animal upon any of these ends does not depend upon its acquired experience, but upon some feature of its innate constitution; and that feature is what we commonly and properly call an instinct, an innate tendency to strive after some end of a particular kind, an innate conative
tendency."

The reactive tendency of my definition differs from Lasurski's Hauptneigung in that it is meant to connote something more than an artificial abstraction, and to refer to specific properties of the organism rather than to the generally inclusive traits of personality that Lasurski has in mind. It differs from McDougall's innate conative tendency in its recognition of the fact that the features of an animal's innate constitution are plastic, and capable of modification by experience. I am also inclined to approach the analysis of behavior from a somewhat different viewpoint than that which is implied in McDougall's statements. In dealing with behavior one is apt to be diverted from the most proximate aspect of the phenomena under consideration by estimating the facts solely with reference to the ultimate needs of the individual or, more usually, of the species. It is somewhat artificial, I believe, to assume that a given sequence of activities is set in operation by outer stimuli acting in conjunction with a need which is more apt to be a product of the biologist's analysis than (in other than a rather mystical sense) dynamically a part of the animal's reactive equipment. The preferable course is to identify individual hungers which are the product of inner,

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physiological events and environmental stimulations. The satisfaction of a given hunger may or may not be conducive to the welfare of either the individual or the speciesmay or may not correspond to a biological need. The total phenomenon with which the behaviorist is concerned in a given case consists of a sequence of events of which the first member is usually, but not always, an external situation. Then come the physiological processes that produce the hunger-impulsion to which the reaction is to be ascribed. The identification of a reactive tendency becomes possible whenever we are able to predict that the operation of a given hunger-impulsion will be directly followed by a series of activities which conform to a known type. The existence and nature of a given hunger is, of course, arrived at by inference; but this does not necessarily call for a departure from a purely objective attitude toward the facts. If, as Watson (4) suggests, the behaviorist may use the term " consciousness " as it is used by other natural scientists, no insuperable difficulty ought to be encountered in the construction of criteria for the identification of a hunger as a relatively independent dynamic unit. Such criteria would take into account the facts concerning the physiology of the sense-organs to which we already have access; and would recognize the various possibilities for experiencing specifically different satisfactions by employing different modes of stimulating the sense-organs. For example, when I entered the laboratory yard this morning carrying a pail of loquats, the coyote leaned against the large meshes of the wire fence that confines him and whined until I scratched his head; the monkeys gave their characteristic food-calls until I gave them loquats; and after the male monkeys had eaten this agreeable addition to their breakfast they sought the females and copulated with them. I did not need to raise a question as to the contents of each animal's state of consciousness to assist me in the identification of the coyote's dog-like hunger for a mild irritation to his cutaneous sense-organs, or the monkeys' hunger for the various stimulations that are derived from eating food, etc. The coyote may or may not have had a mental picture of the head-scratching or the monkeys of the loquats that usually appear these days whenever I put my hand into the little tin pail; but I am quite sure that these animals were clamoring in response to the particular hungers that I have come to recognize,

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and that their behavior, the types of which I could have predicted before I entered the yard, was the expression of definite organic properties. The essential points of the above discussion are these: (1) The essential factors concerned in behavior phenomena are (a) the action of physiological processes usually operating in conjunction with environmental forces, in the production of (b) hungers which impel the individual to manifest (c) activities, the particular types or modes of which are to be ascribed to (d) specific organic properties (reactive tendencies). (2) These reactive tendencies are most conveniently classified with reference to the individual hungers that bring them to expression. (3) The term " reactive tendency," according to my definition, is meant to designate something more specific than an inclination to direct activity toward one of a limited number of general ends, and to include both the innate and the acquired features of an individual's reactive mechanism. CONCLUSIONS At least two, and possibly three, different kinds of hunger, or needs of individual satisfaction, normally impel the macaque toward the manifestation of sexual behavior, viz., hunger for sexual satisfaction, hunger for escape from danger and, possibly, hunger for access to an enemy. Homosexual beh'avior is normally an expression of tendencies which come to expression even when opportunities for heterosexual intercourse are present. Sexually immature male monkeys appear to be normally impelled toward homosexual behavior by sexual hunger. The fact that homosexual tendencies come to less frequent expression in the mature than in the immature male suggests the possibility that in their native habitat these animals may wholly abandon homosexual behavior (except as a defensive measure), on arriving at sexual maturity. Homosexual behavior is of relatively frequent occurrence in the female when she is threatened by another female, but it is rarely manifested in response to sexual hunger. Masturbation does not seem to occur under normal conditions. The macaque of both sexes is apt to display sexual excitement in the presence of friendly or harmless non-primates. It is possible that the homosexual behavior of young males is

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of the same biological significance as their mock combats. It is clearly of value as a defensive measure in both sexes. Homosexual alliances between mature and immature males may possess a defensive value for immature males, since it insures the assistance of an adult defender in the event of an attack.
REFERENCES
(1) FREUD, SIGMUND. Drei Abhandlungen zur Sexualtheorie. Vienna. 1905. (2) LASURSKI, A. Ueber das Studium der Individualist. Pddagogische Mono1912. graphien, XIV Band, Leipzig. (3) MCDOUGALL, WM. The Sources and direction of psycho-physical energy 1913. American Journal of Insanity, vol. 69, no. 5, p. 865. (4) WATSON, JOHN B. Psychology as the behaviorist views it. Psychologica 1913. Review, vol. 20, no. 2, pp. 158-178.